Orthopteran Communities in the Conifer-Broadleaved Woodland Zone of the Russian Far East

Home , Bryodemella, Dianemobius, Mecostethus, Oecanthus pellucens, Pteronemobius, Ruspolia nitidula

Eur. J. Entomol. 105: 673–680, 2008 http://www.eje.cz/scripts/viewabstract.php?abstract=1384 ISSN 1210-5759 (print), 1802-8829 (online)

Orthopteran communities in the conifer-broadleaved woodland zone of the Russian Far East

THOMAS FARTMANN, MARTIN BEHRENS and HOLGER LORITZ*

University of Münster, Institute of Landscape Ecology, Department of Community Ecology, Robert-Koch-Str. 26, D-48149 Münster, Germany; e-mail: [email protected]

Key words. Orthoptera, cricket, grasshopper, community ecology, disturbance, grassland, woodland zone, Lazovsky Reserve, Russian Far East, habitat heterogeneity, habitat specifity, Palaearctic

Abstract. We investigate orthopteran communities in the natural landscape of the Russian Far East and compare the habitat requirements of the species with those of the same or closely related species found in the largely agricultural landscape of central Europe. The study area is the 1,200 km2 Lazovsky State Nature Reserve (Primorsky region, southern Russian Far East) 200 km east of Vladi- vostok in the southern spurs of the Sikhote-Alin Mountains (134°E/43°N). The abundance of Orthoptera was recorded in August and September 2001 based on the number present in 20 randomly placed 1 m² quadrates per site. For each plot (i) the number of species of Orthoptera, (ii) absolute species abundance and (iii) fifteen environmental parameters characterising habitat structure and microclimate were recorded. Canonical correspondence analysis (CCA) was used first to determine whether the Orthoptera occur in ecol- ogically coherent groups, and second, to assess their association with habitat characteristics. In addition, the number of species and individuals in natural and semi-natural habitats were compared using a t test. A total of 899 individuals of 31 different species were captured, with numbers ranging between 2 and 13 species per plot. Species diversity was higher in semi-natural habitats than natural habitats. There was a similar but non-significant pattern in species density. Ordination analysis indicated four orthopteran communities, which were clearly separable along a moisture and vegetation density gradient. The natural sites in the woodland area of the Lazovsky Zapovednik are characterized by species-poor and low-density orthopteran assemblages compared to the semi-natural sites. But, the natural sites have a higher diversity of habitat specialists. Our findings corroborate the hypothesis that intermediate habitat disturbance levels support particularly species-rich animal communities at high densities. Under such regimes, orthopterans presumably mostly profit from the high diversity in plant species, which generates great structural and microclimatic heterogeneity.

INTRODUCTION America, where different aspects of rangeland grass- While natural forests in Europe were to a great extent hopper communities have been studied in detail (e.g. transformed by man into agricultural land and settlement, Kemp et al., 1990; Kemp, 1992a, b; Fielding & Brusven, huge areas of the East Palaearctic are still forested (New- 1993a, b, 1995; Joern, 2004, 2005). Most community ell, 2004; Yan & Shugart, 2005) and thus are important studies in the Palaearctic are for central Europe and dry reference areas for the study of temperate woodland land- and semi-dry grassland habitats (e.g. Fartmann, 1997; scapes. The Far East is one of the three biodiversity hot- Behrens & Fartmann, 2004). Information on the Asian spots in Russia (Venevsky & Venevskaia, 2005) and a part of the Palaearctic is restricted to biogeographic data centre of diversity and endemism of Orthoptera in Eura- (Stebaev et al., 1989; Sergeev, 1998) and detailed studies sia (Sergeev, 1998). of orthopteran assemblages are lacking. Since woodlands Their taxonomy and distribution are well studied, and are usually not considered to be an orthopteran habitat the ease with which they can be sampled and their func- (Theuerkauf & Rouys, 2006) and old forests are rare in tional importance make Orthoptera suitable subjects for central Europe, little is known about habitat selection and ecological and biogeographical studies (Sergeev, 1997; community structure of Orthoptera in natural woodland Lockwood & Sergeev, 2000). Habitat selection in areas in the Palaearctic. Orthoptera is based on a complex combination of dif- We therefore investigated orthopteran communities in ferent and often interrelated environmental factors. Of the natural landscape of the Russian Far East and com- these parameters, the microclimate at oviposition sites, pared the results with observations from the human and which is often affected by vegetation structure, plays a agriculturally dominated landscape of central Europe, crucial role (Uvarov, 1977; Willott & Hassall, 1998). because many taxa occur throughout the Palaearctic (Ser- Sergeev (1997) stressed the suitability of orthopteran geev, 1992, 1997). Hence, orthopteran assemblages in the communities for ecological and biogeographical investi- Lazovsky State Nature Reserve (Primorsky region, Rus- gations. In recent decades many such studies have been sian Far East) were studied to (i) determine their species done in the northern hemisphere. Especially in North composition and abundance in different natural and semi- natural habitats, (ii) analyse orthopteran habitat require-

* Present address: Helmholtz-Centre for Environmental Research Leipzig-Halle UFZ, Department of Community Ecology, Theodor-Lieser-Str. 4, D-06120 Halle, Germany.

673 At the coast winters are warmer (average January temperature: –11°C) and summers cooler (average August temperature: 17°C) (Semenchenko, 2003). Sampling of Orthoptera Sampling was carried out on 18 plots representing all the typical orthopteran habitats of the Lazovsky Zapovednik, except floodplains, which were studied by Specht (2004). Nine natural (coastal dunes, semi-dry coastal grasslands, swamps) and nine semi-natural habitats (fallows dominated by Artemisia spp. and meadows) were investigated. The area of the plots was > 2,000–10,000 m2 with a homogenous vegetation structure at every site. Orthopteran densities were recorded in box quadrats (Gardiner et al., 2005) of a total area of 20 m2. From 29/08–15/09/2001 one sample was taken on each plot: The mobile 1 × 1 m (1 m2) and 80 cm high quadrat was randomly placed at twenty different points. Sampling was done in sun- shine at temperatures > 20°C, between 10:00 a.m. and 5:00 p.m. Except for the small Nemobiinae species, which live hidden under stones or in litter on the ground, sampling provided reli- able quantitative data. Most of the specimens were determined in the field and then released. Individuals that could not be identified in the field (Tetrix spp., some Chorthippus spp.) and voucher specimens of each species were collected and identified later. For determination the keys of Bey-Bienko & Mishchenko (1951a, b) and Storozhenko (1986) were used. Nomenclature is based on Storozhenko (1986) and for species that also occur in Europe on Heller et al. (1998). Fig. 1. Study area and location of Lazovsky Zapovednik. Habitat structure ments in relation to habitat structure and microclimate For each plot we measured/estimated fifteen environmental and (iii) compare orthopteran habitat preferences there parameters: inclination, exposure, heights of one (minimum) up to three different vegetation layers (e.g. turf – tall grass – Artem- and in Europe where the same or closely related species isia) and % cover of the following habitat components: total occur. vegetation, field layers, Cyperaceae, Poaceae, herbs, mosses, lit- MATERIAL AND METHODS ter, bare soil, stones and hollows (in swamps). Data analysis Study area Canonical correspondence analysis (CCA) (using CANOCO The study area is located in the Primorsky region (southern 4.51; ter Braak & Šmilauer, 2002), a direct gradient ordination Russian Far East, Fig. 1). The landscape is formed by the technique, was used to determine the organization of orthop- Sikhote-Alin Mountains, stretching from the southwest to the teran species into distinct communities and the relations northeast, parallel to the coastline (average altitude about between habitat structure and species composition (Fielding & 1,000 m a. s. l.). Woodland covers 80% of the Primorsky region Brusven, 1993b, 1995; Palmer, 1993; Szövényi, 2002; Torrusio – taiga in the north, conifer-broadleaved woodlands in the south et al., 2002). Environmental and species data were log arithmi- (Newell, 2004). cally transformed [y’ = ln (y + 1)] to obtain approximately The 1,200 km2 sized Lazovsky State Nature Reserve (i.e. normal distributions and homogenous variances. Species of Lazovsky “Zapovednik”, the official Russian category for this Orthoptera that occurred only on one plot and/or of which protected area) is situated about 200 km east of Vladivostok in < 10 specimens in total were found were not included in the data the southern spurs of the Sikhote-Alin Mountains (134°E/43°N). set (Table 1). Inclination and exposure were not used as vari- It mainly consists of woodlands dominated by Mongolian oak ables in the CCA because only two plots were slightly inclined (Quercus mongolica) with an admixture of Korean pine (Pinus (< 5°); cover (%) of bare soil and stones made up one variable; koraiensis) and various other tree species. The species richness out of the three field layer heights the maximum vegetation of the Zapovednik is impressive (1212 species of vascular height was used in CCA. The statistical validity of the ordina- plants, 57 mammals and 318 birds), with many rare and highly tion was tested using a Monte Carlo permutation test (null endangered species including the Amur tiger (Panthera tigris model: 9,999 unrestricted permutations). This was carried out altaica) (Chochrjakow & Schochrin, 2002; Newell, 2004). Open for every environmental variable and all canonical axes (i.e. the habitats are very rare, and include natural coastal dunes and complete model). Only significant variables were included step- swamps, parts of the floodplains, screes and mountain peaks, wise in the model, and at each step only the variable that anthropogenic meadows in woodland-clearings near the three explained most of the remaining error variance (manual-forward ranger camps and a few set-aside fields at the reserve border. selection of CANOCO) was chosen. Non-significant variables A monsoon climate with warm, humid summers and cold, dry were those that explained little of the additional variance at the winters is characteristic of the study area. The average annual time they could be added to the model. They also may intercor- precipitation is 750–850 mm, decreasing from the coast inland. relate with other environmental variables (Storch et al., 2003); Due to a greater influence of continental climate the mean tem- which was examined using Spearman’s rank correlation perature inland is –20°C in January and 20°C in July–August. analysis.

674 TABLE 1. The 14 most common orthopteran species on the 18 plots in order of their fidelity. Species that occurred only on one plot and/or with < 10 specimens in total are not included (see Data analysis). Distribution in Europe: species that also occur in Europe are indicated by an “+”. Exlusiveness: species that are restricted to natural (n) and semi-natural (sn) habitat types are indicated. Fidelity (no. of Sum of Density Distribution Exclusive- Species Abbreviation occupied plots) specimens (ind./10 m²) Europe ness Phaneroptera falcata (Poda, 1761) Ph.fal 13 54 1.50 + . Polionemobius taprobanensis (Walker, 1869) Po.tap 12 270 7.50 . . Tetrix japonica (Bolivar, I., 1897) Te.jap 9 23 0.64 . . Oecanthus longicaudus Matsumura, 1904 Oe.lon 8 45 1.25 . sn Ruspolia nitidula (Scopoli, 1786) Ru.nit 5 31 0.86 + n Chorthippus maritimus Mishchenko, 1951 Ch.mar 4 12 0.33 + n Chorthippus hammarstroemi (Miram, 1908) Ch.ham 4 64 1.78 . sn Chorthippus schmidti (Ikonnikov, 1913) Ch.sch 4 13 0.36 . . Omocestus haemorrhoidalis (Charpentier, 1825) Om.hae 4 23 0.64 + n Dianemobius fascipes nigrofasciatus Di.fas 3 39 1.08 sn (Matsumura, 1904) Mecostethus parapleurus (Hagenbach, 1822) Me.par 3 16 0.44 + sn Teleogryllus infernalis (Saussure, 1877) Te.inf 3 13 0.36 . . Oxya maritima Mishchenko, 1951 Ox.mar 2 36 1.00 . n Pteronemobius nitidus (Bolivar, I., 1901) Pt.nit 1 116 3.22 . n

T tests (using SPSS 11.5) were used to assess significant dif- Orthopteran assemblages ferences in orthopteran density and species number for the plots Five of the ten environmental variables significantly at the natural and semi-natural sites. Prior to the analyses, variables were tested for normal distribution using Kolmogorov- contributed to the CCA ordination model. The model Smirnov test. explained 60% of the variance in the number of species and 96% of the variance in the species-environment rela- RESULTS tion (total inertia: 3.08, sum of all canonical eigenvalues: Species richness and abundance 1.93; Monte Carlo test: F = 4.04, P d 0.001). Each of the four canonical axes was highly correlated with one par- A total of 31 species (9 Tettigoniidae, 5 Gryllidae, ticular environmental variable, facilitating the ecological 1 Tetrigidae and 16 Acrididae) and a sum of 899 speci- interpretation of each axis (see Table 2 for details). All mens were captured. Species number ranged from 2 to 13 non-significant variables intercorrelated with other envi- per plot. Phaneroptera falcata and Polionemobius tapro- ronmental variables (Table 3) and explained < 10% addi- banensis were the most widespread species occurring in tional variance at the stage when they would have been 13 (72%) and 12 (67%) of the plots, respectively (Table included in the model. 1). The total number of species was higher at semi-natural Based on the first two canonical axes four distinct com- (mean values ± SE: 9.11 ± 0.75) than natural sites (5.11 ± munites can be separated (Fig. 2a): Cover of hollows 0.82) (t test, t = –3.582, df = 16, P = 0.01). Similarly strongly correlated with the first canonical axis. Note that orthopteran density [individuals (ind.)/10 m², excluding there is no “real” environmental moisture gradient under- Nemobiinae] was higher at semi-natural (mean values ± lying this ordination pattern, because there are only two SE: 17.44 ± 4.21) than natural sites (8.89 ± 2.15). How- plots (swamps) with hollows. Along the first canonical ever, the difference was not significant (t test, t = –1.812, axis species are divided into two distinct main groups: df = 16, P = 0.089). In general there was a positive rela- Taxa that were recorded only in swamps (natural habitat) tionship between species richness and overall orthopteran and those only in mesic to dry habitats (semi-natural and density (Y = 0.735x + 3.628, R2 = 0.27, P < 0.05). Of the natural habitats). The swamps have a species-poor semi-natural sites, young fallows (N = 4) had the highest orthopteran community. However, two species, Oxya species numbers (9.75 ± 1.49) and densities (28.50 ± 5.52 maritima and Pteronemobius nitidus, were restricted to ind./10 m²). In contrast to the total number of species, that the wetlands. of abundant species restricted to one of the habitat types Based on the arrangement of species along the second was more or less the same (Table 1). Five species canonical axis it is possible to further discriminate occurred exclusively in natural (swamps and dunes: between of the orthopteran communities in the remaining Chorthippus maritimus, Omocestus haemorrhoidalis, habitats. The axis is negatively correlated with herb Oxya maritima, Pteronemobius nitidus and Ruspolia niti- cover. The strongest positive correlations (in order of dulus) and four in semi-natural habitats (meadows and arrangement) with herb cover are for Mecostethus para- abandoned fields: Chorthippus hammarstroemi, Diane- pleurus, Chorthippus hammarstroemi, Oecanthus longi- mobius fascipes nigrofasciatus, Mecostethus parapleurus caudus and Dianemobius fasciatus. They form a second and Oecanthus longicaudus).

675 TABLE 2. Summary of CCA for 14 orthopteran species (N = 755 specimens) and five significant environmental variables (F-values of Monte Carlo test, ***P < 0.001, *P < 0.05). Environmental Axis 1 2 3 4 F Eigenvalue 1.00 0.38 0.28 0.20 Species-environment correlations 1.00 0.85 0.88 0.76 Variance explained (%): Species data 32.5 12.4 8.9 6.4 Species-environment relation 51.8 19.7 14.0 10.5 Linear correlation with cover (%) of: Hollows 1.00 0.00 0.02 0.017.69*** Herbs –0.21 –0.71 0.26 –0.15 2.10* Poaceae –0.46 0.20 0.71 –0.25 2.15* Field layers 0.04 –0.23 0.48 –0.83 2.19* Cyperaceae 0.42 0.54 –0.06 –0.42 2.15* community found in herb-rich fallows and meadows (semi-natural habitat). Clearly separated from all other species and strongly negatively correlated with herb cover is Chorthippus maritimus. On the most extreme dune sites (natural habitat) with sparse vegetation and short swards Chorthippus maritimus is typical of the third species-poor community. Sporadically a few other species occur at low density. Also, but to a lower extent Omocestus haemorrhoidalis and Ruspolia nitidula are negatively correlated with herb cover. Both species occur syntopically in coastal semi-dry grasslands (natural habitat) when they form part of another orthopteran community. Further insights in the relations between species of Fig. 2. Biplots for the axes of canonical correspondence Orthoptera and vegetation structure are indicated by the analysis (CCA): a – first vs. second axis; b – third vs. fourth third and fourth axes (Fig. 2b). The folllowing species are axis. Significant environmental/ground cover variables (arrows) positively correlated with the cover of field layers and and position of orthopteran species (crosses). Abbreviation of Poaceae: Ruspolia nitidula, Phaneroptera falcata and species names (see Table 1). Two variables (hollows, herbs), Polionemobius taprobanensis. R. nitidula was most abun- which do not correlate with axis three and four (see Table 2), dant in Poaceae-rich and dense coastal grasslands. P. fal- i. e. their vectors were close to zero, are not shown in b. cata and P. taprobanensis were found in nearly all habitats, except the coastal dunes with extensive areas of A group of four species (Chorthippus hammarstroemi, bare ground and swamps. Teleogryllus infernalis, Dianemobius fascipes and Chor- thippus maritimus), which are further from the origin of

TABLE 3. Correlation matrix of Spearman’s rank correlation analyses (N = 18; ***P < 0.001, **P < 0.01, *P < 0.05). Ground cover (%) Environmental Vegetation Bare soil/ Field variables height (cm) Hollows Litter Moss Cyperaceae Poaceae Herbs stones layers Ground coverage Vegetation total 0.31 0.39 –0.87*** 0.44 –0.18 0.39 0.37 0.24 0.83*** Field layers 0.56* 0.11 –0.71** 0.67** –0.37 0.13 0.53* 0.49* Herbs 0.48* –0.20 –0.27 0.37 –0.33 –0.20 –0.04 Poaceae 0.23 0.01 –0.33 0.37 –0.07 –0.27 Cyperaceae –0.23 0.39 –0.53 –0.03 0.40 Moss –0.41 0.51* –0.23 –0.36 Litter 0.23 –0.12 –0.49* Bare soil and stones –0.14 –0.45 Hollows 0.14

676 the axis, is strongly negatively correlated with cover of density. High orthopteran densities are often the result of field layers and Poaceae. C. hammarstroemi, D. fascipes a trade-off between optimal microclimatic conditions on and T. infernalis reach their highest densities in fallows the one hand and sufficient food and low predation pres- with bare soil. The first two are restricted to these sites. sure on the other (Fielding & Brusven, 1992; Gottschalk, 1996; Fartmann & Mattes, 1997; Behrens & Fartmann, DISCUSSION 2004). Of crucial importance for Orthoptera abundance Species richness and abundance are the microclimatic conditions during egg and larval According to Stebaev et al. (1989) many of the development (Ingrisch, 1979, 1980). In this period most observed orthopteran species are associated with the species benefit from high temperatures (van Wingerden et southern border of the forest zone and limited to the al., 1991a). In the sparse vegetation of the coastal dunes Pacific Ocean districts of the Russian Far East with a microclimate may be favourable, but food shortage and monsoon climate. Numerous species are concentrated in easy access for avian predators seem to result in a low the study area, in regions of conifer-broadleaved forest orthopteran density. An increase in vegetation density and forest-steppe in the south of the Russian Far East results in lower temperatures in the egg habitat, mostly (Stebaev et al., 1989; Sergeev, 1992). located near the soil surface (van Wingerden et al., The natural sites in the woodland area of the Lazovsky 1991a). Although, the dense stands of the old set-aside Zapovednik are characterized by species-poor and low- fields should provide enough food and enemy-free space, density orthopteran assemblages compared to the semi- the microclimate seems to be unfavourable for many natural sites. At the semi-natural sites the highest densi- orthopteran species. Because the young open Artemisia ties and species numbers were found on young fallows. fallows in the Lazovsky Zapovednik provide the best The young set-aside fields studied were characterized by combination of requirements, the orthopteran abundance the highest structural heterogeneity. there is the highest. What are the reasons for a high orthopteran diversity at Orthopteran assemblages the semi-natural sites? Several studies (Fartmann & Presence and assemblage of orthopteran species are dis- Mattes, 1997; Kruess & Tscharntke, 2002; Gebeyehu & tinctly different among habitat types. This depends on Samways, 2003) indicate that orthopteran species rich- orthopteran habitat preference, which is determined by ness is highest at sites subject to intermediate disturbance species adaptation to habitat structure, microclimate and (e.g. grazing) and high structural heterogeneity. Joern disturbance intensity (Joern, 1982; Fielding & Brusven, (2005) found that grasshopper species richness in grass- 1995; Samways, 1997; Szövényi, 2002). The distribution land is positively correlated with plant species richness pattern of eurytopic and stenotopic species differ – due to and heterogeneity of vegetation structure, and negatively their specific habitat requirements with the latter species with vegetation height and grass biomass. Sites subject to characteristic of different habitat types in the Lazovsky an intermediate level of disturbance (like young fallows) Zapovednik. Where possible, the following compares the have more plant species (Grime 1973a, b) and therefore a present results (Fig. 2, Table 1) with ecological observa- greater structural heterogeneity. Thus, greater resources tions from Europe. are available for the coexistence of more orthopteran spe- The eurytopic and thermophilous species Phaneroptera cies (Dennis et al., 1998). In our view oviposition sites falcata and Polionemobius taprobanensis are not and food resources are the most important, especially the restricted to one orthopteran community and occur in presence of bare ground for oviposition can be a limiting both natural and semi-natural habitats in the study area. factor. Many, or even the majority of orthopteran species However, their habitat requirements differ. The terrico- lay their eggs in the ground and prefer bare or sparsely lous, flightless cricket P. tabrobanensis is found in high covered ground for oviposition (Richards & Waloff, densities in short-turf vegetation. It is a thermophilous 1954). In this study the number of orthopteran species species distributed throughout the Indo-Malayan region exclusive to natural and semi-natural sites is similar (5 vs. and the southern Far East (Schmidt 1999). The 4 species). Including the orthopteran data of Specht phytophilous/arbusticolous and very mobile (capable of (2004) for the floodplains of the Lazovsky Zapovednik flying) bush-cricket P. falcata occurs on those habitats at and thus all the typical natural orthopteran habitats in the low densities. It prefers tall herbaceous vegetation and study area, five further species are restricted to natural has a Transpalaearctic distribution (Detzel, 1998a). habitats [Bryodemella tuberculata (Fabricius, 1775), Dia- The following groups of species have a higher habitat nemobius csikii (Bolivar, I., 1901), Eirenephilus longi- specifity and form four distinct orthopteran communities: pennis (Shiraki, 1910), Oedaleus infernalis Saussure, 1884 and Tetrix tenuicornis (Sahlberg, 1893)]. Orthop- 1. Community of coastal dunes with bare ground teran assemblages of the natural sites of the Lazovsky The geophilous Chorthippus maritimus, a far eastern Zapovednik are species-poor, but have a greater diversity sibling species of C. biguttulus (Linnaeus, 1758) (Krivo- of highly specialised species. lutskaya, 1997), occurs in the study area only in natural In accordance with the results of Joern (2005) there is a and dynamic habitats with extensive areas of bare ground positive relationship between overall Orthopteran density or stones, such as coastal dunes, floodplains or screes and species richness, indicating that there are at least (own observation). In the floodplains of the Lazovsky some parameters that promote both species diversity and Zapovednik it typically co-occurs with Bryodemella

677 tuberculata and Eirenephilus longipennis (Specht, 2004). cies P. heydenii (Fischer, 1853) with a Holomediterra- C. biguttulus, a Eurosiberian and common species in cen- nean origin (Kiechle, 1998) and a Mediterranean- tral Europe, shows different habitat requirements: it is -southwest Asian distribution (Baur et al., 2006). phytophilous and eurytopic with a preference for P. heydenii is characterised as hygrophilous, terricolous semi-dry grassland (Fartmann, 1997; Behrens & Fart- and thermophilous; typical habitats in central Europe are mann, 2004). south-facing lakeshores, swamps or wet meadows 2. Community of semi-dry coastal grassland (Kiechle, 1998; Winterholler & Bierwirth, 2003). Ruspolia nitidula, a Palaeotropic/Mediterranean species CONCLUSIONS is thermophilous, phytophilous/graminicolous and espe- The majority of Orthoptera in the boreal and temperate cially the immature stages are hygrophilous (Detzel, forest zones of the Palaearctic are associated with open 1998b). The restriction of this bush-cricket to coastal habitats (Sergeev, 1992, 1997). These grassland habitats grassland with a moderate maritime climate in the often are created and maintained by disturbance Lazovsky Zapovednik seems to result from these adapta- (Theuerkauf & Rouys, 2006), which is especially impor- tions. R. nitidula prefers vertical structures and in the 2 tant at the microhabitat scale. At this level, disturbance study area it is most abundant (up to 7 individuals/10 m ) creates the microclimatic conditions and microhabitats in ecotones of dense, tall-grass stands, which possibly necessary for the survival of eggs and larval stages. This offer a suitable microclimate and structure. applies to Orthoptera and other thermophilous insects The stenotopic Omocestus haemorrhoidalis, a Transpa- (e.g. Lepidoptera: Fartmann, 2004, 2006). As for the laearctic species, has similar vegetation structure prefer- whole temperate zone of the Palaearctic (Birks, 2005; ences in the Lazovsky Zapovednik and in central Europe. Mitchell, 2005), closed forests are the natural vegetation It is geo-/phytophilous, graminicolous, xero- and thermo- of most parts of the Lazovsky Zapovednik. In the absence philous. Microhabitat preferences in the Russian Far East of man, open habitats are restricted to sites with extreme correspond to those observed by Fartmann (1997) in environmental conditions. Only sites that are highly dis- northeast Germany: open, short-turf semi-dry grassland turbed (dunes, floodplains), too dry (dunes, screes) or wet with structural heterogeneity and bare soil. (swamps) and too cold (mountain tops) are free of trees, 3. Community of herb-rich meadows and fallows all the intermediate sites are covered with woodland. Due Mecostethus parapleurus and Oecanthus longicaudus to the origin of the landscape one would expect that the are herbicolous and thermophilous species of the herb- hot spots of orthopteran diversity and density will be sites rich fallows and meadows in the study area. M. para- created by natural disturbance. But, this is not the case as pleurus is distributed along the southern boundary of the semi-natural sites subject to anthropogenic disturbance nemoral woodland zone from the Atlantic Ocean to (recently set-aside fields) have the highest species num- Japan; in central Europe it is a thermophilous species, bers and abundance. The orthopteran communities of the which prefers wet meadows (Detzel, 1998c). The habitat natural habitats are not as species-rich as the anthropo- requirements of O. longicaudus seem to resemble those of genic sites, but there are more stenotopic species the Mediterranean O. pellucens (Scopoli, 1763) in central restricted to those habitats. Europe: According to Detzel (1998d) O. pellucens is a Comparable observations exist for central Europe; but typical species of tall forb-rich plant communities. disturbance intensity as well as the spatial and temporal Chorthippus hammarstroemi is an eastern Palaearctic, pattern in the cultural European landscape are very dif- meso-xerophilous steppe species (Sergeev, 1997) that is ferent from a natural landscape. Human activity in central abundant in anthropogenic habitats, such as fields (Ser- Europe creates either extensive and continuously high geev, 1998). In the Lazovsky Zapovednik the cricket Dia- disturbance (e.g. intensive cropland) or approximately nemobius fascipes nigrofasciatus occurs syntopically with stable conditions with low disturbance frequency (e.g. C. hammarstroemi exclusively in recently set-aside fields, high forests with long cutting intervals), which both lead but reaches its highest density in open, dry fallows with to habitat loss for insects of open habitats. Natural, much bare soil/stones. Hence, this terricolous cricket is dynamic disturbance patterns and low-intensity anthropo- considered to be thermo- and xerophilous. On the Kuril genic management have positive effects on habitats and Islands it colonises soils heated by volcanic activity, close populations of stenotopic insects (Sergeev, 1998; Di to solfataras and fumaroles (Krivolutskaya, 1997). It is Giulio et al., 2001), but today such practices are rare in widely distributed in the temperate region of East Asia; central Europe (Fartmann, 2006). Hence, our findings in on the Japanese Islands the southern range limit is about the natural landscape of the Russian Far East confirm a 30°N (Masaki, 1996). strategy of nature conservation that has recently attracted interest in central Europe. That is, the conservation and 4. Community of swamps restoration of natural landscape dynamic/disturbance The hygrophilous and phytophilous/graminicolous (Finck et al., 1998) and support of anthropo-zoogenic Oxya maritima (Catantopinae) occurs together with the management with low intensity (van Wingerden et al., terricolous cricket Pteronemobius nitidus in the coastal 1991b; Wallis De Vries & Raemakers, 2001; Theuerkauf swamps. Masaki & Oyama (1963) describe P. nitidus as a & Rouys, 2006) for a lasting conservation of insect biodi- typical species of paddy fields in North Japan. The habitat versity. requirements seem to resemble those of the sibling spe-

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